Rev. Biol. Trop.

, 49(2): 697-702, 2001

www.ucr.ac.cr www.ots.ac.cr www.ots.duke.edu

Within-tree distribution of Ecdytolopha torticornis (Lepidoptera:

Tortricidae) oviposition on macadamia nuts
Helga Blanco-Metzler1, Allan D.Watt2 and Derek Cosens3
1

2
3

Centro de Investigacin en Proteccin de Cultivos-Estacin Experimental Fabio Baudrit Moreno,

Universidad de Costa Rica, Facultad de Agronoma, San Jos, Costa Rica, Fax (506) 433-9086,
E-mail: hblanco@cariari.ucr.ac.cr
Banchory Research Station, Hill of Brathens, Glassel, Banchory AB31 4BY, Scotland, UK
E-mail: adw@wpo.nerc.ac.uk
Institute of Cell, Animal and Population Biology, The University of Edinburgh, Ashworth Laboratories,
West Mains Road, Edimburgh EH9 3JT, Scotland, UK E. Mail: Derek.Cosens@ed.ac.uk

Recibido 22-II-2000. Corregido 27-IX-2000. Aceptado 17-I-2000.

Abstract: Vertical distribution of eggs of the macadamia nutborer Ecdytolopha torticornis Meyrick (Lepidoptera:
Tortricidae) and its preference of oviposition sites within and between macadamia cultivars were studied in Turrialba,
Cartago, Costa Rica, in 1992 (N = 6 939). E. torticornis eggs were found throughout the foliar parts of the tree, but
fewer eggs were laid in the crown top than in the mid or lower crown. Differences in the horizontal distribution of
the eggs were not significant, albeit more eggs were found in the outer positions. The numbers of eggs found within
the crowns of different clones were similar, implying that the nutborer has no preference for a particular cultivar.
Key words: Ecdytolopha torticornis, oviposition, sampling, macadamia, Costa Rica.

Since it was first recorded in Costa Rica (F.

Lara 1987 unpublished) the nutborer
Ecdytolopha torticornis Meyrick (Lepidoptera:
Tortricidae), has been responsible for an
increase in damage to the nuts of Macadamia
integrifolia Maiden and Betche (Proteaceae)
from a loss in yield of 16 % in 1987 (Lara
1987), to 28% in 1990 (C.E Mass and L.F.
Campos 1990 unpublished), and 39 % in 1992
(Blanco-Metzler et al. 1994).
The E. torticornis female lays its eggs singly
on the middle zone of macadamia nuts and,
usually, in the narrow space between adjacent
nuts within a cluster. Upon emergence the larva
bores into the nut and feeds on the husk, if the nut
shell has not hardened the larva will bore

through, and feed on the kernel (Blanco et al.

1993). The biology of E. torticornis closely
resembles that of the koa seedworm
Cryptophlebia illepida (Butler) (Lepidoptera:
Tortricidae), and the litchi fruit moth, C.
ombrodelta (Lower) (Jones and Caprio 1992). E.
torticornis moths have been reported to be poor
fliers (Chamberlain 1989) living in the tree
canopy (Blanco-Metzler 1994), and since the
larvae feed inside the nut, they are limited in their
ability to disperse. Thus, the distribution of
larvae and hence nut damage are largely
determined by egg distribution.
The oviposition behaviour of the nutborer
within the host plant has provided useful and
necessary information for the development of

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sampling methods. A survey of total egg mass is

one of the methods used in determining pest
abundance in many horticultural crops as
Diaphania nitidalis (Stoll) (Lepidoptera:
Pyralidae) in cucumber (Blanco-Metzler 1983);
Helicoverpa zea (Boddie) (Lepidoptera:
Noctuidae) in maize (Hoffmann et al. 1991), and
in forest pests such as the gypsy moth, Lymantria
dispar (L.) (Lepidoptera: Lymantriidae) in birch
trees (Higashiura 1989, Thorpe and Ridgway
1992) or the spruce budworm, Choristoneura
fumiferana (Clem.) (Lepidoptera: Tortricidae) in
spruce trees (Lysyk 1990). The aim of this study
was to investigate the within-tree distribution of
the nutborer by recording its oviposition sites,
and to determine whether moths showed
ovipositional preferences between cultivars.

MATERIALS AND METHODS

The study site was located at Oriente Farm,
Turrialba, Costa Rica, at elevations ranging from
620 to 700 m above sea level. The vertical
distribution of E. torticornis eggs was studied in
a mixed stand planted in 1978 of four of the most
commonly planted cultivars in Costa Rica:
Keahou (HAES 246), Kau (HAES 344), Kakea
(HAES 508) and Keaau (HAES 660) with a
height ranging from 4.5 m to 6 m. The study
period was from July to December 1992. The
nuts were examined in the laboratory (23 C,
80 % RH) at CATIE (Tropical Agronomical
Research and Higher Education Centre,
Turrialba), and the following variables were
evaluated: the number of eggs per nut; the
number of fertile and infertile eggs and empty
chorions; and the number of nutborer holes per
nut. The number of holes was included as an
indication of multiple, successful ovipositions.
Some nuts were found to have holes but no
chorions were present, perhaps because they had
been disloged by rain, blown away by wind, or
eaten by predators. The same trees were sampled
every month. Trees were divided into three

crown levels (Fig. 1): I, crown top, 3.2 m from

the ground to the top of the tree crown position
1); II, mid-crown, 1.6 - 3.2 m from the ground
(positions 2 and 3); III, lower-crown, 0 - 1.6 m
from the ground (positions 4 and 5). The height
of level III corresponded to that of the first
author. Levels II and III were each divided into
two positions: positions 3 and 5 = nuts sampled
in an area up to 1 m horizontally from the tree
trunk; positions 2 and 4 = nuts sampled in an
area up to 1 m horizontally from the end of the
branches. Twenty nuts (over 0.8 cm in diameter)
per crown level were collected at random; ten
nuts per position for crown levels II and III.
The data were submitted to Analysis of
Variance and T tests (LSD) (Anonymous,
1985). The experimental design used was a
split split plot where the tree was the big plot,
the combination of levels and positions was
the split plot and the time (6 months, July to
December) was the split split plot. Data were
transformed by sqrt (x + 0.5) as many nuts
were free of eggs. Nuts were collected every
month from five trees per clone, 20 trees in
total. In the statistical analysis, crown level I
was compared with levels II and III together;
level II versus level III; and the inner sampling
positions (3 and 5) were compared with the
outer sampling positions (2 and 4).

Fig. 1. Diagrammatic representation of the sampling

positions and levels used in determining the vertical
distribution of the nutborers eggs on macadamia.

INTERNATIONAL JOURNAL OF TROPICAL BIOLOGY AND CONSERVATION

RESULTS
A total of 6 939 nuts were inspected. Newly
laid eggs were frequently observed on nuts
already bearing eggs or bore-holes. The number
of eggs laid per nut varied from cero to eight
with a mean of 0.8 (n = 201 nuts). No significant
differences were found in the number of eggs per
cultivar (F = 0.69; d.f. = 3; p = 0.57).
Eggs were found on macadamia trees at all
heights; however, they were significantly more
abundant in crown levels II and III than in level
I (p < 0.001). The mean number of eggs per nut
in top level I was 0.14; in mid level II: 0.29;
and in bottom level III: 0.30. There was no
significant difference between the number of
eggs found on nuts from the lower two levels
(F = 0.22; d.f. = 1; p = 0.64); nor between the
mean number of eggs found on nuts on the
inner and outer parts of the branches (F = 2.0;
d.f. = 1; p = 0.16) (Table 1). While fewer eggs
were found on the inner sampling positions (3
and 5) than on the outer sampling positions (2
and 4), no statistical difference was found
between the number of eggs in either positions
for levels II and III (F = 0.22; g.l. = 1; p = 0.64).

699

There was a significant difference (p <

0.0001) in the number of eggs per nut over the
six months of sampling (Table 2). The number
of eggs per nut was highest from July to
October, decreased in November (0.2) and
dropped steeply to a minimum (0.06) in
December (Fig. 2). The decline in the number
of eggs per nut paralleled the decline in nut
production, but not the decline in the number of
damaged nuts which occurred approximately
one month earlier (Fig 2).

TABLE 2
Numbers of Ecdytolopha torticornis eggs laid per
macadamia nut over time, 1992*
Month
(1992)

Number of
nuts sampled

Mean number of
eggs per nut

July
August
September
October
November
December

1149
1000
1190
1200
1200
1200

0.34 a
0.28 a
0.29 a
0.34 a
0.20 b
0.06 c

* p = 0.05

Crown
Level Position

Number of nuts
sampled

Mean number of
eggs per nut

Level Position

Level

Position

2233

2223

0.14 b

0.14 b

II

2
3

2377

1198
1179

0.29 a

0.30 a
0.28 a

4
5

2329

1157
1172

0.30 a

III
* p = 0.05

0.32 a
0.28 a

40

0.4

30

0.3

20

0.2

10

0.1

0.0

Mean number of eggs per nut

Numbers of Ecdytolopha torticornis eggs laid per

macadamia nut by crown position*

Means: Number of falen nuts

and % of damaged nuts

TABLE 1

May Jun Jul Aug Sep Oct Nov Dic

Month

Fig. 2. The mean number of fallen nuts (divided by 10;

solid bar); the mean percentage of damaged nuts (hatched
bar); and the mean number of eggs per month (line graph).

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DISCUSSION
Oviposition site selection is one of the most
important aspects of habitat selection in insects.
It varies with insect species and is closely related
to offspring survival. The observed habit of
laying eggs on nuts already containing eggs or on
damaged nuts (Blanco-Metzler et. al. 1993)
might give the larvae the advantage of escaping
from parasitoids and predators, or prevent them
from being dislodged by the rain. BlancoMetzler (1994) reported that it took a larva
approximately fifty minutes from emerging from
the egg to completely bore through the husk.
Therefore, if an entrance to the nut has already
been made, the first instar larva is more likely to
evade detrimental environmental factors and
increase its survivorship. Conversely, the
females' habit of laying eggs on damaged nuts
may lead to intra-specific larval competition for
food and/or space. Moreover, larval mortality
might increase due to early nut-drop (BlancoMetzler et. al. 1992; Blanco-Metzler 1994).
Sinclair (1979) observed that Cryptophlebia
ombrodelta, also tended to lay its eggs
preferentially on damaged macadamia nuts; he
concluded that this behaviour lowered the
chances of survival of the larvae in dense
populations due to cannibalism and early nutfall. However, cannibalism was not observed in
E. torticornis: yet early nut-fall was observed
and concomitant enhanced larval predation
(Blanco-Metzler et al. in preparation).
Oviposition in insects has been reported to
be influenced by physical properties of the host
such as shape of the trees and nuts, colour of
leaves and nuts, surface texture of fruits
(Prokopy and Bush 1973) and chemical
stimulants of the host (Renwick and Radke 1983;
Hedin and McCarty 1990). Since no differences
were found in the number of eggs between
cultivars it is unlikely that during oviposition
females were influenced by cultivar variation in
shape, size, and colour of the nuts. Thus, the
differences in nut damage for cultivars 246, 344,
508 and 660 (C.E. Mass and L.F. Campos 1990;
Blanco-Metzler, et. al. 1992; Blanco-Metzler
1994) might rather reflect differences in larval

survival resulting from cultivar differences in

food quality due to variation in secondary plant
compounds and husk toughness. Indeed, damage
was found to decrease with nut maturation
because as the nut dries it provides food of a
poorer quality and quantity.
The tendency of finding most eggs in the outer bottom level of the tree crown, followed by
the outer middle level, could be because nut production is highest in this part of the tree (Diego
Prez, personal communication), and therefore,
the moths lay their eggs where food abundance is
highest to ensure the best possible food supply for
their offspring. Also, since E. torticornis has been
reported as probably being a poor flyer (Chamberlain 1989), the adults reach this part of the tree
more readily from the weeds where they are believed to feed. Thus during the period of oviposition, they do not have to fly far to find their host
and lay their eggs. In Oriente Farm weed control
is limited as they have found that weeding increases root fungi problems and soil erosion (Diego
Prez, personal communication).
There are three probable reasons that could
explain the differences in the number of eggs per
nut found over time. First, during the period
September to November nut production was
high and females found plenty of oviposition
sites: as time passed and fewer nuts (oviposition
sites) were available and, thus, were thus harder
to find. Secondly, a reduction in the number of
nuts restricts the available larval food resource
and in turn the larval population, hence fewer
adults will eventually emerge. Thirdly, the
seasonal increase in the abundance of parasitoids
(Blanco-Metzler 1994) is also likely to cause a
decline in the size of the nutborer population.
Although Cryptophlebia females show no
preference for laying eggs at different heights in
the tree crowns (Jones and Tome 1992) E.
torticornis females do prefer to oviposit on nuts
growing within the first three metres of the
ground. We reaffirm comments by Southwood
(1978) and Horn (1988) that the identification of
host characteristics relevant to oviposition
behaviour and insect distribution (both within
and between host plants) is important for the
design of sampling programmes, the forecasting

INTENATIONAL JOURNAL OF TROPICAL BIOLOGY AND CONSERVATION

of insect damage, and for assessment of the

effectiveness of control programmes.
AKNOWLEDGMENTS
This research was supported by the British
Council, AID-ROCAP, the Tropical Agronomical Research and Higher Education Centre (CATIE) and Macadamia de Costa Rica.

RESUMEN
Se determin la distribucin vertical de los huevos del
barrenador de la nuez de macadamia Ecdytolopha torticornis
Meyrick (Lepidoptera: Tortricidae) y los sitios de preferencia
de oviposicin en los rboles y entre clones de macadamia.
Se detect la presencia de huevos de E. torticornis en todo el
rbol, sin embargo, se encontr un menor nmero de huevos
en la parte alta de la corona que en la parte media e inferior.
La diferencia en la distribucin horizontal de los huevos fue
no significativa, a pesar de encontrarse un mayor nmero de
huevos en las posiciones externas. El nmero de huevos entre clones fue similar, sugiriendo que la polilla del barrenador
no tiene preferencias de oviposicin entre clones.

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